Research Article |
Corresponding author: Hua Peng ( hpeng@mail.kib.ac.cn ) Corresponding author: Hang Sun ( sunhang@mail.kib.ac.cn ) Academic editor: Hanno Schaefer
© 2021 Yi Yang, Hua Peng, Hang Sun.
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Yang Y, Peng H, Sun H (2021) Taxonomic revision of Sageretia (Rhamnaceae) from China I: identities of S. lucida, S. thea var. cordiformis and S. yunlongensis, with the description of a new species S. ellipsoidea. PhytoKeys 179: 13-28. https://doi.org/10.3897/phytokeys.179.64750
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A taxonomic revision of Sageretia lucida, S. thea var. cordiformis and S. yunlongensis in China is presented. Sageretia lucida is revised in terms of morphological characters (habit, branchlet color, phyllotaxis and rachis length), distribution, habitat, and phenology; S. thea var. cordiformis is raised to S. cordiformis; and S. yunlongensis is excluded from the genus Sageretia and reduced to the synonym of Rhamnus nigricans. Furthermore, a new species, S. ellipsoidea, is erected based on the paratype collections of S. lucida. The new species morphologically differs from S. lucida in having reddish brown branchlets, opposite or subopposite phyllotaxis, shorter rachises, and flowering in spring or early summer. S. ellipsoidea is factually closest to S. hamosa as they share similar woody-vine habit and larger fruit size, and fruiting in winter, whereas the former can be easily recognized based on its smaller leaf blades, fewer lateral veins, shorter rachises, and ellipsoidal or elliptic-ovoid fruits.
Granite mountain, limestone mountain, Rhamnus, woody vine
Sageretia Brongn., the mock buckthorn genus of Rhamnaceae, contains ca. 35 species (
According to Flora of China (FOC;
We have been studying the taxonomy, molecular phylogeny and biogeography of Sageretia since 2014, especially the members in China. A new species, the tropical Asian origin, and three strongly supported clades matching morphological and distributional divergences, had been reported in Sageretia in our previous studies (
The protologues of all published names and molecularly phylogenetic studies of Sageretia, were carefully reviewed and collated. Specimens or digital specimen images from 26 herbaria, including A, CDBI, CSH, CZH, CSFI, E, FJSI, GXMG, HHBG, HITBC, IBK, IBSC, IMDY, K, KUN, LBG, LD, P, PE, SWFC, S, SYS, SZG, UC, US and WU (abbreviations follow
Sageretia yunlongensis G.S.Fan & L.L.Deng, Sida 16(3): 477, f. 1. 1995. syn. nov. Type: China. Yunnan: Yunlong County, 1300 m, 26 Oct 1987, Expedition Team 161 (holotype SWFC!)
China. Yunnan: “Beyendjing medium inter Tschuhsiung (Tsuyung) et Yungbei”, 1800 m, 15 May 1915, Hand.-Mazz. 6311 (holotype WU!; isotypes A [00051422], K [K000729152]).
Type specimens of Rhamnus nigricans and its synonym A isotype of R. nigricans (K [K000729152]) B holotype of S. yunlongensis (SWFC). A obtained from GBIF (https://www.gbif.org/), B photographed by Y. Yang.
Evergreen vines, shrubs or small trees up to 6 m tall, dioecious. Young branches yellowish-brown pubescent; old branches scattered with tuberculate lenticels. Leaves alternate; stipules caducous; petioles 1.2–2.5 cm; leaf blades papery or thickly papery, ovate, oblong to broadly elliptic, 5–16 × 3–7 cm, abaxially puberulent or only on veins, adaxially usually glabrous, lateral veins 5–7 pairs, prominent abaxially, impressed adaxially, base rounded to subcordate, margin densely cartilaginous serrulate, apex acuminate to shortly caudate. Inflorescences axillary, spicate or paniculate, rachises up to 10 cm, puberulent. Flowers unisexual, 5-merous; pedicels 1–2 mm; sepals triangular; petals clawed. Drupes subglobose or globose, ca. 6 mm in diam., turning purple-black at maturity; pyrenes 2–3, asymmetrical, abaxially with a margined furrow extending over 3/4 of length.
Flowering from May to July; ripe fruits from October to December.
The species is distributed in southwestern China (Yunnan; Fig.
Although the genera Rhamnus and Sageretia are similar in morphology, they distinctly differ in characters of inflorescence (fascicled, cymose racemes, or cymose panicles in Rhamnus vs. spikes or spicate panicles in Sageretia) and fruits (basally persistent discoid calyx tube in Rhamnus vs. persistent reflexed calyx or remaining inconspicuous disk in Sageretia). The type collection of S. yunlongensis, Expedition Team 161, has branched cymose panicles and fruits basally covered with discoid calyx tube, suggesting it belongs to Rhamnus rather than Sageretia. In fact, S. yunlongensis (Fig.
China. Yunnan: Weishan Yi and Hui Autonomous County, 1500 m, 2012, Weishan Expedition Team 5329271259 (IMDY); Yongde County, 1830 m, 9 Jul 2006, E.D. Liu 170 (KUN); Shuangbai County, 1670 m, 15 Apr 1957, W.Q. Yin 747 (KUN, LBG, PE); Lushui County, 1544 m, 12 May 2005, Gaoligong Shan Biodiversity Survey 23964 (E).
S. thea var. cordiformis Y.L.Chen & P.K.Chou in Bull. Bot. Lab. North-East. Forest. Inst. 5: 74. 1979. Basionym.
China. Yunnan: Xishuangbanna, Mengla County, 730 m, 28 Dec 1958, W.T. Wang 10496 (holotype KUN [1207932]; isotype KUN [1207933]).
Type specimens of Sageretia cordifolia, S. cordiformis and S. thea A, B isotypes of S. cordifolia (A: P [P01818867]; B: P [06765093]) C holotype of S. cordiformis (KUN [1207932]) D isotype of S. cordiformis (KUN [1207933]) E, F isotypes of S. thea (E: S [S11-12914]; F: LD [1749752]). A, B and E, F obtained from GBIF, C, D from NOI (https://noi.link/).
Evergreen shrubs. Branches usually alternate, armed, glabrous to puberulent; second- to fourth-year branches brown. Leaves alternate or subopposite; petioles 1–3 mm, leaf blades leathery, shiny, glabrous, cordate to ovate-orbicular, 1–3 × 0.8–2 cm, lateral veins 2–3 pairs, flat abaxially, impressed adaxially, base cordate or subcordate, margin crenate, apex obtuse or rounded. Inflorescences spicate or spicate-paniculate; rachises 1.5–5 (–10) cm. Flowers sessile; sepals triangular-ovate; petals clawed; stamens as long as petals. Drupes subglobose, ca. 5–6 mm in diam., from green to red and finally turn black, base with persistently reflexed calyx; pyrenes 2–3, emarginate at both ends, asymmetrical.
Flowering in September; ripe fruits from December to January of the following year.
The species is distributed in China (southwestern Yunnan; Fig.
In Flora Yunnanica,
Comparison of habitat and morphology among Sageretia cordifolia, S. cordiformis, and S. thea based on field observation, herbarium collections, and photo information obtained from CFH and PPBC.
S. cordifolia | S. cordiformis | S. thea | |
---|---|---|---|
Habitat | unknown | limestone mountains | hills or mountains |
Branch color | gray | gray-brown to brownish | brownish |
Petiole length | 4–8 mm | 1–3 mm | 2–7 mm |
Leaf blade shape | ovate-oblong to ovate-lanceolate, base cordate, apex acute to caudate-acuminate | cordate or ovate-orbicular, base cordate or subcordate, apex obtuse or rounded | elliptic, oblong, or ovate-elliptic, rarely ovate or nearly orbicular, base rounded or subcordate, apex acute, obtuse, or rounded |
Leaf blade size | 3.5–6 × 1.5–3 cm | 1–3 × 0.8–2 cm | 2–4.5 × 0.7–2.5 cm |
Leaf texture | leathery | leathery | paper |
Lateral veins | 3–5 pairs | 2–3 pairs | 3–5 (–7) pairs |
Rachis | 12–13 cm | 1.5–5 (–10) cm | 2–12 cm |
Based on specimen examination and our field observations, S. thea var. cordiformis and the type variety (Fig.
Laos. Pakson, 1200 m, Nov 1938, E. Poilane 28562 (holotype P [01818865]; isotypes P [P01818866, P01818867, P06765093]).
China. Yunnan: Mengla County, 25 Sept 1961, Y.H. Li 3588 (KUN); 1200 m, 24 Nov 1975, Y.H. Li 20033 (HITBC); 1000 m, 9 Sept 1959, S.C. Pei 10046 (HITBC); 10 Sept 2004, S.S. Zhou 2084 (PE); ca. 1000 m, 21 Dec 2015, Y. Yang & Z. Chen xsbn03 (KUN); ca. 1000 m, 24 Dec 2016, Y. Yang & L.S. Qian OYY001 (KUN).
China. Guangdong: “Ying Tak, Taai Tsan, Wan Tong Shan” (Yingde City, Taizhen Town, Wentang Mountain), 17 Oct 1926, W.T. Tsang & K.C. Wong 2718 (holotype IBSC [0404901]; isotype SYS [SYS00086833]); 18 Oct 1926, W.T. Tsang & K.C. Wong 2723 (paratype IBSC [0404896]); 6 Oct 1926, W.T. Tsang & K.C. Wong 2479 (paratype SYS [SYS00086832]).
Holotype of Sageretia ellipsoidea (IBSC [0404901]). Image obtained from CVH (https://www.cvh.ac.cn/).
Similar to S. hamosa (Wall.) Brongn., but differs in having smaller leaves (5–12 × 2.5–4 cm in S. ellipsoidea vs. 8–15 (–25) × 3.5–6 (–7) cm in S. hamosa), less lateral veins (5–7 pairs vs. 7–11 pairs), shorter rachises (1–3 (–10) cm vs. 6–20 (–25) cm), and ellipsoidal or elliptic-ovoid fruits (vs. subglobose or globose in S. hamosa).
Woody vines. Branches opposite or subopposite, glabrous; first-year branches green, sometimes with hard-straight spine opposite to leaf, second- to fourth-year branches reddish brown. Leaves opposite or subopposite; petioles 8–15 mm, leaf blades leathery, ovate-oblong, oblong to elliptic, 5–12 × 2.5–4 cm, lateral veins 5–7 pairs, prominent abaxially, impressed adaxially, base rounded, margin crenate, apex obtuse to shortly acuminate. Inflorescences usually axillary spicate, rarely spicate-paniculate; rachises 1–3 (–10) cm. Flowers subsessile, white to yellowish white; sepals triangular-ovate, ca. 1.5 mm, apex acute; petals clawed; stamens as long as petals. Drupes ellipsoidal or elliptic-ovoid, 10–12 × 5–7 mm, green, turning to orange-red, claret and finally purple-black, base with inconspicuous disk remains; pyrenes 1–2, emarginate at both ends, asymmetrical.
Flowering from April to July, ripe fruits from November to January of next-year.
This species is named for its ellipsoidal or elliptic-ovoid drupes which are different from other Sageretia species (subglobose or globose).
The species is currently found in southern China (Fujian, Guangdong, Guangxi, Hainan; Fig.
When he erected the species Sageretia lucida,
Besides, the samples of “Sageretia lucida” in
Comparison of habitat, morphology and phenology (florescence) among Sageretia ellipsoidea, S. hamosa and S. lucida based on field observation, herbarium collections, and photo information obtained from CFH and PPBC.
S. ellipsoidea | S. hamosa | S. lucida | |
---|---|---|---|
Habitat | granite mountains | non-limestone hills or mountains | limestone mountains |
Habit | woody vines | wood vines | shrubs |
Phyllotaxis | opposite or subopposite | alternate or subopposite | alternate |
Branch color | reddish brown | reddish brown to brown | gray to dark gray |
Petiole length | 8–15 mm | 8–15 (–20) mm | 6–10 mm |
Leaf blade shape | ovate-oblong, oblong or elliptic, base rounded, apex obtuse to shortly acuminate | usually oblong or narrowly elliptic, base usually rounded, sometimes cordate, apex caudate-acuminate to shortly acuminate | ovate-oblong to oblong, base subrounded to rounded, apex acuminate |
Leaf blade size | 5–12 × 2.5–4 cm | 8–15 (–25) × 3.5–7 cm | 5–10 × 2.5–4 cm |
Lateral veins | 5–7 pairs | 7–11 pairs | 4–6 pairs |
Rachis | 1–3 (–10) cm | 6–20 (–25) cm | 5–10 cm |
Fruits | ellipsoidal or elliptic-ovoid | globose or subglobose | unknown |
Florescence | April to July | July to August | November |
China. Fujian: Minhou County, 4 Oct 2014, B. Chen & D.M. Jin CSH12700 (CSH); Nanjing County, 400 m, 19 Nov 1963, Xiamen Univ. Coll. Team 805 (PE); Pinghe County, 600 m, 23 Feb 1980, G.S. He 0475 (FJSI); Zhao’an County, 950 m, 16 Mar 2015, X.F. Zeng ZXF19893 (CZH). Guangxi: Jinxiu Yao Autonomous County, 200 m, 6 Apr 1982, Dayao Shan Expedition Team 13973 (IBSC); Shangsi County, Shiwandashan, 370–390 m, 14 Nov 2011, D.X. Nong et al. FC2011061 (GXMG). Guangdong: Dinghu District, 8 Nov 1963, G.Q. Ding & G.L. Shi 1132 (IBSC); Fengkai County, 15 Nov 1980, G.Q. Ding & G.L. Shi 6652 (CDBI); Ruyuan Yao Autonomous County, 17 Aug 1935, S.K. Lau 23948 (IBK); Huaiji County, 500 m, 26 Oct 1958, Y.G. Liu 2707 (HHBG); Yangchun City, 6 Nov 1935, C. Wang 38672 (IBK, PE); Chao’an District, 900 m, 18 Oct 2009, X.F. Zeng ZXF8404 (CZH); Conghua District, 600 m, 4 Dec 1958, L. Deng 8836 (IBK); Shenzhen City, 300–350 m, 20 Sept 2006, G.D. Wang et al. 6474 (SZG); Boluo County, 444.6 m, 1 Apr 2019, Y. Yang OYY00121 (KUN). Hainan: Baoting Li and Miao Autonomous County, 6 May 1935, F.C. How 72211 (IBK); Changjiang Li Autonomous County, 7 Jun 1934, H.Y. Liang 64162 (IBK); Baisha Li Autonomous County, 29 Apr 1936, S.K. Lau 26548 (IBK); Lingshui Li Autonomous County, 21 Oct 1956, L. Deng 2785 (KUN).
China. Guangdong: “Ying Tak, Taai Tsan, Chung Tung” (Yingde City, Taizhen Town, Zhongdong Village), 24 Nov 1926, W.T. Tsang & K.C. Wong 3260 (holotype UC [319815]; isotypes A [00051501], SYS [SYS00095840], US [00094394]).
Shrubs up to 3 m. Branches alternate, glabrous; second- to fourth-year branches gray to dark gray. Leaves alternate; petioles 6–10 mm, leaf blades leathery, ovate-oblong to oblong, 5–10 × 2.5–4 cm, lateral veins 4–6 pairs, prominent abaxially, impressed adaxially, base sub-rounded to rounded, margin serrulate, apex acuminate. Inflorescences axillary spicate; rachis 5–10 cm. Flowers sessile; sepals triangular-ovate, 1.2–1.5 mm long; petals clawed; stamens as long as petals. Fruits unknown.
Flowering in November; fruits unknown, probably ripening from April to May of the next year.
The species is endemic to Yingde, Guangdong, China (Fig.
Sageretia lucida is closest to S. henryi Drumm. & Sprague in morphology and sharing similar limestone habitats and flowering in autumn. Thus,
China. Guangdong: Yingde City, Zhongdong Village, 13 Nov 1926, W.T. Tsang & K.C. Wong 3114 (SYS).
We thank Dr. Xin-xin Zhu (College of Life Sciences, Xinyang Normal University) and Mr. Jian Lin for providing field photos of Sageretia thea and S. ellipsoidea, respectively. We are grateful to Mrs. Yun-xiao Liu (South China Botanical Garden, CAS) for the line drawings. We also thank the curators of IBSC, KUN, PE and SYS, and Mr. Kai-wen Jiang (Southwest Forestry University) for their assistance with specimen examination. This study is jointly supported by the Second Tibetan Plateau Scientific Expedition and Research (STEP) program (Grant No. 2019QZKK0502) and National Natural Science Foundation of China (Grant No. 31560064).